化感——外来入侵植物的“Novel Weapons”

综述了与外来植物入侵相关的理论假说。并主要介绍了从化感角度提出的“N ove lW eapons”假说(NW假说)。某些外来入侵植物能产生一些化学物质。这些物质可以发挥较强的化感作用,或者成为植物和土壤微生物之间相互作用的调节者。由于不同区域的植物群落共同进化的轨迹不同,被入侵群落的植物对这些化学物质缺乏适应性,因此,这些物质成为了外来种入侵的利器。而对外来种发源地的群落来说,由于长期的共存关系,这些物质对其伴生植物的影响相对较小。NW假说为化感作为外来植物入侵的一种机制奠定了理论基础,为外来入侵植物的防治提供了理论依据,但作为一种较新的理论假说,还需要更多的实验研究加以支持。     

植物光合产物分配及其影响因子研究进展

植物光合产物分配受环境因子和生物因子的共同影响。为增进对植物对全球变化响应的理解, 从植物个体水平与群落/生态系统水平综述了植物光合产物分配的影响因子与影响机理的最新研究进展。植物个体在光照增强及受水分和养分胁迫时, 会将光合产物更多地分配到根系; CO₂浓度升高对植物光合产物分配的影响受土壤氮素的制约, 植物受氮素胁迫时, CO₂浓度升高会促进光合产物更多地分配到根系; 反之, 对植物光合产物分配没有影响。植物群落/生态系统的光合产物分配对环境因子的响应不敏感; 光合产物向根系的分配比例随其生长阶段逐渐降低。功能平衡假说、源汇关系假说和相关生长关系假说分别从环境因子、个体发育和环境因子与个体发育协同作用方面阐述了植物光合产物分配的影响机理。在此基础上,指出了未来拟重点加强的研究方向: 1)生态系统尺度的光合产物向呼吸部分的分配研究; 2)地下净初级生产力(belowground net primary productivity, BNPP)研究; 3)温室和野外条件下及幼苗和成熟林光合产物分配对环境因子响应的比较研究; 4)生态系统尺度的多因子控制试验; 5)整合环境因子和个体发育对植物光合产物分配格局的影响研究。     

群落组成及物种间相互作用对外来植物入侵的影响

外来植物入侵已成为严重的环境和社会问题,了解外来植物的入侵机制是有效控制其入侵的前提。生物阻抗假说认为,入侵地本地植物群落中的许多生物因子及生物过程能够抵御外来植物入侵。但关于群落抵抗外来种入侵的主要机制,目前还没有确定的结论。本文综述了群落中物种功能特征的多样性以及与外来种功能特征的相似度、植物与动物、植物与植物以及植物与土壤微生物间的相互作用等因素对外来植物入侵的影响,以及以前研究存在的不足。未来研究应该注重不同条件下植物与植物间的相互作用;不同竞争强度下,植物与食草动物的相互作用;植物、动物及土壤微生物三者之间的相互作用对外来植物入侵的影响。这些研究不仅能够丰富和完善入侵生态学理论,而且对于预测外来植物未来的扩散范围,合理有效地管理生态系统,防止外来植物入侵,保护本地生物多样性具有重要的实际意义。     

放牧格局和生境资源对矮嵩草（Kobresia humilis）分株生物量分配和补偿性生长的影响

为研究密集型克隆植物对放牧扰动和生境资源变化的生物量分配和补偿生长响应特性,验证克隆植物的觅食模型和3个有关植物个体补偿反应的假说(①放牧优化假说,grazing optimization hypothesis,GOH;②反应连续谱假说,continuum of responses hypothesis,CRH;③增长率模型,growth rate model,GRM),在具有不同放牧利用格局和土壤养分水平的高寒矮嵩草草甸(Ⅰ.畜圈草地:重度放牧、资源丰富;Ⅱ.牧道草地:中度放牧、资源贫乏;Ⅲ.封育草地:不放牧、资源贫乏)中通过设置扣笼/无扣笼样方对其建群种矮嵩草(Kobresia humilis)进行了研究.结果表明不论当年解除家畜放牧与否,春季采摘率越高(畜圈草地),分株生物量向生长的投入越少,向贮藏器官的投入越多.繁殖分配在中度采摘下最高(牧道草地),扣笼内外分株各部分的生物量分配无差异.矮嵩草分株在中度采摘×资源贫乏条件下产生了超补偿响应,在重度采摘×资源丰富条件下为等量补偿,重度采摘导致分株密度显著减少.生物量分配格局与觅食模型的预测不符.补偿生长响应特性证实了GOH和GRM的预测,但与CRH的预测不符.这说明在研究地区放牧扰动格局对克隆植物矮嵩草分株的生物量分配和补偿生长具有重要影响,适度放牧利用更利于引起超补偿,而重度利用可能会对该种群的长期保持产生不利影响.     

森林生态系统中植食性昆虫与寄主的互作机制、假说与证据

围绕"多样性稳定性"假说、"联合抗性假说"、"生长势假说"、"胁迫假说"、以及下调、上调和推拉等机制与假说提出的背景与实验验证的证据,力图辨析其概念以及它们之间的相互关系。作者认为,多样性-稳定性机制关注森林生态系统的功能,是基于群落甚至景观层次。多样性条件下的联合抗性机制和联合易感性应属于稳定性中的抵抗力范畴。联合抗性机制的主要基础是基于资源集中假说和天敌假说,这些观点在种群层次上更易理解;上调力和下调力机制是以食物网底部的资源与顶端的天敌来探讨这种互作关系。因此,资源集中与上调力有着对应关系,而天敌假说只是下调力机制中的一个层面而已。植物生长势假说和植物胁迫假说力图从植物个体或种的群体的生长状态出发解析植食性动物的对寄主的选择趋势。上述有关植食性昆虫与寄主互作的机制、假说与证据是基于不同的层面提出的,因而在解析研究目标时,由于基本面的差异有可能会得出不同的结论。以近年来的研究进展和研究成果为依据有针对性地阐述这些理论对森林有害生物生态调控技术的指导作用,其中,联合抗性和联合易感性理论对指导森林有害生物生态控制具有更直接的指导作用。进一步提出了相应的亟待解决的科学问题。     

土壤类型对短葶飞蓬灯盏乙素和咖啡酸酯积累的影响

在相同栽培管理条件下,分别以黑色石灰土、红色石灰土、红壤、黄红壤、紫色土和腐殖土为生长介质对短葶飞蓬（Erigeron breviscapus）进行了人工栽培试验。结果显示,生长在不同土壤类型的短葶飞蓬植株体内总咖啡酸酯和灯盏乙素含量有显著的差异。红色石灰土、紫色土及黄红壤生长的植株总咖啡酸酯和灯盏乙素含量最高,黑色石灰土和腐殖土的最低。相同土类的不同亚类土壤生长植株间2种次生代谢产物含量的差异程度,会超过不同土类间的。植株总咖啡酸酯和灯盏乙素含量与土壤P和K含量没有显著相关关系,但与土壤速效N含量及植株含N量则都有极显著负相关关系。不同土壤生长植株叶片的最大光能转换效率（Fv／Fm）值与灯盏乙素和总咖啡酸酯含量都表现出极显著负相关关系。土壤类型对短葶飞蓬次生代谢影响规律与碳素／营养平衡假说（CNB）及资源获得假说（HA）的预测一致。     

植物种群表观竞争机制及其研究进展

植物表观竞争既影响群落结构和组成,也与生态系统功能密切相关。目前,国内尚缺乏对植物表观竞争的实验研究和机制探索。对植物表观竞争概念和研究进展进行了系统梳理和阶段性总结,分析了草食动物为媒介的表观竞争主要通过食物和生境介导植物竞争过程,以及ELP-采食者关系对表观竞争的影响;植物参与的表观竞争通常与资源竞争共同作用;AM真菌为媒介的表观竞争不仅受到宿主植物菌根依赖性的影响,还与AM真菌对资源的需求和环境资源供给状况有密切关系。未来研究既要注重实验设计的合理性,也要考虑测定方法和评价参数的有效性;重视植物群落和生态系统水平上表观竞争的功能研究。这不仅能够为植物群落结构和物种共存提供机制性解释,而且能够丰富经典的植物竞争理论和多营养级相互作用的食物网理论。     

Effects of soil nitrogen levels on growth and defense of the native and introduced genotypes of alligator weed

土壤氮水平对喜旱莲子草原产地和引入地基因型生长和防御的影响 同种植物生长在资源丰富生境中的个体,其防御水平被认为低于生长在资源匮乏生境中的个体。然而,生境的养分水平如何影响植物的诱导抗性和耐受性,以及这种影响在入侵植物的原产地和引入 地种群间是否存在差异,目前均知之甚少。本研究以入侵植物喜旱莲子草(Alternanthera philoxeroides)的原产地阿根廷和引入地美国的基因型为研究对象设计同质园实验,以探究土壤氮水平对植物的生长、组成和诱导性[莲草直胸跳甲(Agasicles hygrophila)取食诱导]化学防御以及耐受性的影响。实验中,我们测定了植物总生物量、伸长速率(生长速率的表征)以及叶片和根系中总碳、总氮和三萜皂苷(化学防御物质)的含量。研究结果显示,植物在低土壤氮水平下表现出较高的组成抗性(植物在低土壤氮水平下的叶片三萜皂苷含量高于其在高土壤氮水平的33%)和耐受性[植物被取食后总生物量下降的程度更低(植物在高土壤氮水平和低土壤氮水平下被取食后总生物量分别下降了24%和15%)],而在高土壤氮水平下表现出较高的诱导抗性(在高土壤氮水平下的植物被取食后叶片三萜皂苷含量与空白对照的植物相比升高了24%)。植物的组成抗性和耐受性与生长速率存在权衡,但诱导抗性与生长速率存在显著的正相关性。此外,引入地基因型在低土壤氮水平下叶片碳含量显著低于原产地基因型(-6%),但这种差异在高土壤氮水平下消失。这些结果表明,土壤氮水平 影响植物对不同防御策略的选择偏好,并且在决定引入地基因型的表现时与植食作用存在交互作用。     

植物群落稀有种维持机制与土壤反馈的研究进展

自Janzen-Connell (J-C)假说提出后半个世纪以来, 生态学家在热带及亚热带森林对该假说开展的大量实证研究表明, 由专性天敌导致的J-C效应所引起的负密度制约是维持森林多样性和决定群落组成的重要驱动力, 该假说成功地解释了热带及亚热带森林的丰富多样性。土壤病原真菌所引起的植物-土壤负反馈是J-C效应最主要的表现形式。然而, 对于植物-土壤负反馈是否能够维持森林群落中的大量稀有种仍然存在许多争议。基于当代物种共存理论的“稀有种优势”假说认为, 只有在满足“可入侵准则” (即物种在稀有时具有种群增加的趋势)的前提下, 稀有种才能在群落中与其他物种长期共存。然而, 当前基于土壤反馈的实验结果与该理论预测相悖, 因此在稀有种的维持机制方面仍存在较大的分歧。本文通过介绍植物-土壤反馈理论, 整合了可能对稀有种维持有较大影响的因素, 包括共生菌根真菌、土壤养分以及植物细根性状等在影响土壤负反馈方面的相关研究, 并对这些因素如何影响群落中物种多度和稀有种在群落中的维持进行了探讨。最后, 我们也从其他角度探讨了一些对稀有种维持的研究。我们认为在未来对稀有种的研究中, 探讨使其长期存续的“优势”和制约其种群扩大的“限制”同等重要, 将当代物种共存理论与新技术、新方法相结合对于探究稀有种的维持机制具有重要的意义, 可为稀有种保护提供理论依据。     

植物群落中物种小尺度空间结构研究

植物群落中, 物种小尺度空间结构影响着种群或群落的动态及有关的生态学过程。植物主要是和它同种或异种的邻近个体相互作用,植物个体周围的局部环境和大空间尺度下群落的平均水平是完全不同的。群落中的许多过程都影响小尺度空间结构的形成和动态,同样,局部空间结构反过来影响着植物的生长、更新和凋亡等重要过程。鉴于目前对小尺度空间结构进行的大量研究以及其重要性,有必要对其研究进展进行适当的总结,以期明确将来进一步的研究方向。该文以此为出发点,首先介绍了植物群落中物种小尺度空间结构产生的6个原因:1)生境的空间异质性;2)植物繁殖体的传播;3)植物之间的相互作用;4)生物环境(动物和微生物)的作用;5)外界干扰的作用;6)多因子综合作用。然后阐述了小尺度空间结构意义及对生物多样性、植物种群遗传学和恢复生态学研究的影响。最后对目前物种小尺度空间结构研究存在的几点问题及将来的研究方向作以下归纳:1)大尺度植被动态的研究应该整合小尺度空间结构的信息;2)不论从生物学还是生态学上来讲, 植被小尺度空间结构的研究应该把植物作为中心,确定适当的尺度和采取合理的空间统计方法;3)充分重视小尺度空间结构在退化生态系统恢复中的应用意义;4)注重从小尺度的局部格局研究入手对群落总体特征进行整合;5)植物群落动态研究中,物种小尺度空间结构与平均场假说相结合的必要性。     

Costs of Defense and a Test of the Carbon-Nutrient Balance and Growth-Differentiation Balance Hypotheses for Two Co-Occurring Classes of Plant Defense

One of the goals of chemical ecology is to assess costs of plant defenses. Intraspecific trade-offs between growth and defense are traditionally viewed in the context of the carbon-nutrient balance hypothesis (CNBH) and the growth-differentiation balance hypothesis (GDBH). Broadly, these hypotheses suggest that growth is limited by deficiencies in carbon or nitrogen while rates of photosynthesis remain unchanged, and the subsequent reduced growth results in the more abundant resource being invested in increased defense (mass-balance based allocation). The GDBH further predicts trade-offs in growth and defense should only be observed when resources are abundant. Most support for these hypotheses comes from work with phenolics. We examined trade-offs related to production of two classes of defenses, saponins (triterpenoids) and flavans (phenolics), in Pentaclethra macroloba (Fabaceae), an abundant tree in Costa Rican wet forests. We quantified physiological costs of plant defenses by measuring photosynthetic parameters (which are often assumed to be stable) in addition to biomass. Pentaclethra macroloba were grown in full sunlight or shade under three levels of nitrogen alone or with conspecific neighbors that could potentially alter nutrient availability via competition or facilitation. Biomass and photosynthesis were not affected by nitrogen or competition for seedlings in full sunlight, but they responded positively to nitrogen in shade-grown plants. The trade-off predicted by the GDBH between growth and metabolite production was only present between flavans and biomass in sun-grown plants (abundant resource conditions). Support was also only partial for the CNBH as flavans declined with nitrogen but saponins increased. This suggests saponin production should be considered in terms of detailed biosynthetic pathway models while phenolic production fits mass-balance based allocation models (such as the CNBH). Contrary to expectations based on the two defense hypotheses, trade-offs were found between defenses and photosynthesis, indicating that studies of plant defenses should include direct measures of physiological responses.     

Resource availability and the trichome defenses of tomato plants

We conducted two experiments to determine how resource availability influenced allocation by tomato (Lycopersicon esculentum) to trichomes, and how different patterns of trichome allocation by plants grown in different resource environments might then influence the behavior of tobacco hornworm (Manduca sexta) caterpillars. In the first experiment we used high and low levels of light and water, and then, using scanning electron microscopy, determined trichome densities on the leaves and stems. We sampled leaves and stems at several places throughout the plant to determine whether there were within-plant differences in allocation to trichomes. The results of the first experiment showed that resource availability influenced allocation to trichome growth. Patterns in high and low-light supported both the growth-differentiation balance hypothesis (GDBH) and the carbon-nutrient balance hypothesis (CNBH). However, the GDBH was not supported by differences among water treatments. Contrary, to predictions of the GDBH, plants with intermediate growth did not have the highest trichome densities, and plants with similar growth differed in trichome density. Possible biological and artifactual explanations are discussed. The first experiment also showed that there was within-plant variation in allocation to trichomes, and that plant resource availability may influence within-plant variation in allocation to trichomes. In the second experiment, we grew plants in high and low-light, and then monitored the behavior of tobacco hornworms on the stems of these plants in the laboratory. This experiment demonstrated that the light environment that tomato plants were grown in influenced the resting behavior of caterpillars. Furthermore, it demonstrated that both glandular and non-glandular trichomes impeded caterpillars from searching for food. Overall, this study indicated that plant resource availability can influence allocation to trichome defenses, and that these differences may affect insect herbivores.     

PHLOROTANNIN ALLOCATION AMONG TISSUES OF NORTHEASTERN PACIFIC KELPS AND ROCKWEEDS

Optimal defense theory (ODT) predicts antiherbivore defensive compounds will be allocated so that the most valuable or most susceptible tissues will be best defended. The growth–differentiation balance hypothesis (GDBH) predicts that defense allocation will be a result of trade-offs between growth and defense. Thus, these two theories predict opposite allocation patterns with respect to “valuable,” actively growing meristematic and reproductive tissues. ODT predicts that meristems and reproductive tissues should have higher defense levels than nonmeristematic vegetative tissues; the GDBH predicts the defense levels of meristems and reproductive tissues will be lower than vegetative tissues. We examined allocation patterns of phlorotannins in 21 species of kelps (Order Laminariales) and rockweeds (Order Fucales) from nine sites on the west coast of the United States to determine if allocation patterns better matched the predictions of ODT or the GDBH and to look for differences in allocation patterns among sites. Within-species differences in phlorotannin levels occurred in 10 of the 21 species examined. Meristems of both kelps and rockweeds had higher phlorotannin levels than nonmeristematic vegetative tissues, consistent with ODT. Phlorotannin levels in reproductive tissues of kelps were higher than vegetative tissues, but levels in reproductive tissues of rockweeds were lower than vegetative tissues, indicating that allocation strategies may follow taxonomic lines. Allocation patterns differed among sites in four of the 16 species collected from more than one site. Differences in allocation patterns among sites were usually changes in the ratios of phlorotannins in well-defended compared to poorly defended tissues, rather than changes in which tissues were well defended or poorly defended. We concluded that environmental variability can have large effects on the concentration of phlorotannins in algae but has limited effects on allocation patterns among tissues.     

Plant chemical defense: monoterpenes and the growth-differentiation balance hypothesis

Recent studies of allocation to defensive chemicals in plants have provided insights into the ecological controls over plant defensive chemicals. Both developmental and ecological studies now suggest that we can understand the factors influencing allocation to defense by examining the relative availability of resources, external needs for chemical defense, and the internal demands for growth that plants face. These studies have also shed light on one of the more popular theories in plant evolutionary ecology, the growth-differentiation balance hypothesis of plant resource allocation.     

Testing the optimal defense theory and the growth-differentiation balance hypothesis in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Two prominent theories proposed to explain patterns of chemical defense expression in plants are the optimal defense theory (ODT) and the growth-differentiation balance hypothesis (GDBH). The ODT predicts that plant parts with high fitness value will be highly defended, and the GDBH predicts that slow growing plant parts will have more resources available for defense and thus will have higher defense levels than faster growing tissues. We examined growth rate, fitness value, and defense protein levels in leaves of a wild and lab ecotype of Arabidopsis thaliana to address whether patterns of defense protein expression in this plant conform to predictions of either the ODT or the GDBH. We divided leaves of A. thaliana into six leaf classes based on three developmental stages: vegetative, bolting, and flowering; with two leaf ages at each stage: young and old. We assessed the fitness value of leaves by determining the impact of the removal of each leaf class on total seed production and germination rates. Although A. thaliana was highly tolerant to defoliation, young leaves were more valuable than old in general, and young leaves on bolting plants were the most valuable leaf class in particular. Young leaves on vegetative plants grew fastest in both ecotypes, while old leaves on bolting and flowering plants grew slowest. Finally, defense levels were assessed in each leaf class by quantifying the constitutive and inducible expression of four defense-related proteins. Expression of guaiacol peroxidase and chitinase activity conformed largely to GDBH predictions. Expression of trypsin inhibitor and polyphenoloxidase activity varied by leaf class and treatment, but conformed to neither GDBH nor ODT predictions.     

Testing the growth-differentiation balance hypothesis: dynamic responses of willows to nutrient availability

Here, the growth-differentiation balance hypothesis (GDBH) was tested by quantifying temporal variation in the relative growth rate (RGR), net assimilation rate (NAR), and phenylpropanoid concentrations of two willow species (Salix sericea and Salix eriocephala) across five fertility levels. Initially, RGR increased and total phenylpropanoids declined (although every individual phenolic did not) as fertility increased, but NAR was unaffected. Subsequently, NAR and phenylpropanoids declined in the low fertility treatment, generating a quadratic response of secondary metabolism across the nutrient gradient. As above- and below-ground growth rates equilibrated, NAR and phenylpropanoids increased in the low fertility treatment, re-establishing a negative linear effect of fertility on secondary metabolism. A transient quadratic response of secondary metabolism is predicted when GDBH is integrated with models of optimal phenotypic plasticity, occurring when low NAR imposes carbon constraints on secondary metabolism in low nutrient environments. Once plants acclimate to nutrient limitation, the equilibrium allocation state is predicted to be a negative correlation between growth and secondary metabolism. Although both willow species generally responded according to GDBH, the complexity observed suggests that prediction of the effects of nutrient availability on secondary metabolism (and other plastic responses) in specific cases requires a priori knowledge of the physiological status of the plant and soil nutrient availability.     

Early ontogenetic patterns in chemical defense in Plantago (Plantaginaceae): genetic variation and trade-offs

Predictions based on the plant age and growth-differentiation balance hypotheses of defense were tested in two congeneric species, Plantago lanceolata and P. major, by quantifying iridoid glycosides, defensive chemicals, in seeds and leaves during the first 6 wk of growth. Concentrations decreased from the seed to 2-wk-old seedling stage in P. lanceolata, but increased during this period in P. major. In both species, levels were similar for 2- and 4-wk-old plants, then significantly increased from 4 to 6 wk. Genetic variation in the ontogeny of iridoid glycoside production was significant in both species at the maternal family level and at the population level. To examine whether allocation costs could explain the low production of iridoid glycosides in seedlings, relationships between growth and defense (iridoid glycosides) were characterized. Growth and defense had a positive or null relationship in all age groups, indicating that there was no trade-off in these plants at any age. This study provides some support for the growth-differentiation balance hypothesis, but offers no support for the plant age hypothesis. Measuring how herbivory affects plant fitness at different ontogenetic stages may shed light on these patterns in Plantago and on the evolution of the ontogeny of defense.     

Benefits of the Carbon‐Nutrient Balance Hypothesis

The carbon‐nutrient balance hypothesis (CNBH) is one of a number of approaches to understanding patterns of resource allocation in plants. Numerous empirical tests of the CNBH's predictions have led to certain key refinements and to the recognition that some of the simplifications inherent in the model limit its utility. However, as long as the model is applied to compounds with large pools, and the biosynthetic pathways of secondary metabolites are considered, the CNBH still serves as a useful guide for ecological research on resource allocation. One of the model's values is that it attempts to explain the plasticity of individuals but does not assume that all responses of individuals are optimal in terms of maximizing fitness.     

Interactive effects of habitat productivity and herbivore pressure on the evolution of anti-herbivore defense in invasive plant populations

The evolution of increased competitive ability (EICA) hypothesis predicts that plants released from natural enemies should evolve to become more invasive through a shift in resource allocation from defense to growth. Resource availability in the environment is widely regarded as a major determinant of defense investment and invasiveness, and thus should be incorporated into the conceptual framework of EICA. Analysis of a simple model from the optimal defense literature demonstrates that, in contrast to the EICA hypothesis, enemy release is neither sufficient nor necessary for evolution of reduced resistance among introduced plants when habitat productivity co-varies. In particular, if the invasive range is more nutrient-poor than the native range, there could be selection for more plant defenses even with enemy release.     

Out of the quagmire of plant defense hypotheses

Several hypotheses, mainly Optimal Defense (OD), Carbon: Nutrient Balance (CNB), Growth Rate (GR), and Growth-Differentiation Balance (GDB), have individually served as frameworks for investigating the patterns of plant defense against herbivores, in particular the pattern of constitutive defense. The predictions and tests of these hypotheses have been problematic for a variety of reasons and have led to considerable confusion about the state of the "theory of plant defense." The primary contribution of the OD hypothesis is that it has served as the main framework for investigation of genotypic expression of plant defense, with the emphasis on allocation cost of defense. The primary contribution of the CNB hypothesis is that it has served as the main framework for investigation of how resources affect phenotypic expression of plant defense, often with studies concerned about allocation cost of defense. The primary contribution of the GR hypothesis is that it explains how intrinsic growth rate of plants shaped evolutionarily by resource availability affects defensive patterns. The primary contribution of the expanded GDB hypothesis is that it recognizes the constant physiological tradeoff between growth and differentiation at the cellular and tissue levels relative to the selective pressures of resource availability, including explicitly taking into account plant tolerance of damage by enemies. A clearer understanding of these hypotheses and what we have learned from investigations that use them can facilitate development of well-designed experiments that address the gaps in our knowledge of plant defense.